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Archipelagos and meta- Matthews, Thomas

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Download date: 27. Sep. 2021 UC Merced Frontiers of Biogeography

Title Archipelagos and meta-archipelagos

Permalink https://escholarship.org/uc/item/6w55v67b

Journal Frontiers of Biogeography, 10(3-4)

Authors Whittaker, Robert J. Fernández-Palacios, José M. Matthews, Thomas J. et al.

Publication Date 2018

DOI 10.21425/F5FBG41470

Supplemental Material https://escholarship.org/uc/item/6w55v67b#supplemental

License CC BY 4.0

eScholarship.org Powered by the California Digital Library University of California 1Perspective 2Archipelagos and meta-archipelagos

3Robert J. Whittaker1,2, *, José M. Fernández-Palacios3, Thomas J. Matthews4,5, François

4Rigal5,6, Kostas A. Triantis7

51School of Geography and the Environment, University of Oxford, Oxford OX1 3QY, UK. 62Center for Macroecology, Evolution and Climate, Natural History Museum of Denmark, 7University of Copenhagen, Copenhagen, Denmark. 83Island Ecology and Biogeography Research Group, Instituto Universitario de Enfermedades 9Tropicales y Salud Pública de Canarias (IUETSPC), Universidad de La Laguna, Tenerife, 10Canary , Spain. 114School of Geography, and Environmental Sciences, and Birmingham Institute of 12Forest Research, University of Birmingham, Birmingham B15 2TT, UK. 135Centre for Ecology, Evolution and Environmental Changes (CE3C)–Azorean Biodiversity 14Group and Universidade dos Açores–Depto. de Ciências Agrárias e Engenharia do Ambiente, 15PT-9700-042, Angra do Heroísmo, Açores, Portugal. 166CNRS - Université de Pau et des Pays de l’Adour – E2S UPPA, Institut des Sciences 17Analytiques et de Physico-Chimie pour l'Environnementet les Materiaux, MIRA, UMR5254, 1864000, PAU, France 197Department of Ecology and Taxonomy, Faculty of Biology, National and Kapodistrian 20University of Athens, Athens GR-15784, Greece. 21

22*Corresponding author. Email: [email protected]

23Abstract 24The term meta- has been in use in cultural studies for some time, to refer 25to certain complex areas in which the boundaries between conventionally 26recognised archipelagos are indistinct, although the concept also carries additional 27connotations. Use of the term in biogeography appears more recent and without effort 28to prescribe its meaning. We outline, from a biogeographical perspective, distinctions 29between meta-archipelagos and archipelagos and those islands not occurring within 30either collective grouping, highlighting that network analysis tools provide metrics for 31formal analytical purposes. 32Keywords: Biogeographical regions, island biogeography, meta-archipelago, 33modularity, network analysis 34Text

35 “…the are an island bridge connecting, in “another way,” North to . 36 This geographical accident gives the entire area, including its continental foci, the character of 37 an archipelago, that is, a discontinuous conjunction… [which] can be seen as an island that

1 1 38 “repeats” itself….Which one, then would be the repeating island, Jamaica, Aruba, Puerto 39 Rico, Miami, Haiti, Recife? Certainly none of the ones that we know. That original, that island 40 at the center, is as impossible to reach as the hypothetical Antilli[a] that reappeared time and 41 again, always fleetingly, in the cosmographers’ charts. This is again because the is 42 a meta-archipelago…” Benítez Rojo and Maraniss (1985, p. 431–432) 43 44The etymology of the word archipelago points to a derivation linked to the Italian arcipelago 45as a name for the Aegean (principal ) (Fig. 1). The wider meaning developed presumably 46since the is replete with large numbers of islands. We use the term today for 47chains, clusters, or collections of islands. Yet in complex island regions such as the Caribbean 48or South-East (, New , ) and parts of the South 49Pacific/, it is often debatable where one archipelago ends and another begins 50(Benítez Rojo and Maraniss 1985, above). This matters in island biogeography as many of 51our analyses are based on data sets structured into archipelagos (e.g., Bunnefeld and 52Phillimore 2012). The rationale for this is that islands configured in isolated geographical 53groups exchange information (i.e., there are movements and exchanges of pollen, spores, 54propagules, individuals, semi-nomadic flocks, and perhaps even nutrients and energy) and 55they do so to a significantly greater degree than they do with any other more distant land- 56masses. 57 In analytical terms, some might argue the islands within such archipelagos to be 58spatially auto-correlated and thus non-independent data points. But this depends on the 59questions being asked. In practice, for some purposes island biologists are engaged in 60studying process and pattern at the intra-island level, while for other purposes it is the inter- 61island patterns within the archipelago (such as the species–area relationship) that are the 62focus of interest and analysis (e.g., Whittaker et al. 2017, Price et al. 2018). Moreover, to 63establish the generality of our models and hypotheses, we often wish to extend our analyses 64to encompass islands belonging to many sets of archipelagos (e.g., Bunnefeld and Phillimore 652012, Norder et al. 2018). This generates a further challenge, which is to determine the 66degree to which nearby archipelagos are truly independent ‘replicates’ as opposed to being 67interconnected by similar levels of information exchange as the islands within our 68archipelagos. As, increasingly, evidence of movement behaviours and of past propagule 69exchange and colonization events encoded in phylogenetic data demonstrates that even quite 70distant archipelagos can and do exchange ‘information’ (e.g., Gillespie et al. 2008, Hembry 71and Balukjian 2016), it is not always straightforward to determine ‘natural units’ for specific 72biogeographical analyses. Answering such a question becomes a matter of quantification and 73determining thresholds that might permit objective determination of where the boundaries 74between archipelagos can be drawn (see Box 1). Box 1. Distinguishing the meta-archipelago from the archipelago: a biogeographical definition A Meta-archipelago is a group of archipelagos that have and continue to exhibit a meaningful level of information exchange (e.g., propagules, colonization events) and within which such exchanges occur substantially more often than with other areas but significantly less than the case within a single archipelago. In cases of large, persistent and well isolated systems, the meta-archipelago may be equivalent to a biogeographical sub-region or perhaps to a biogeographical network, but the concept may also be applied to groups of entities within smaller, impermanent and less isolated systems, such as constellations of habitat islands.

2 2 76 77Figure 1. A map of the Aegean, by Nicolaum Visscher (16491702), published around 1681, 78illustrating a complex island region in which it is challenging to decide on archipelago or 79meta-archipelago membership from a biological perspective. Sourced from Wikimedia 80Commons, wherein it is stated that this is an image from the digital and/or physical 81collections of the Koninklijke Bibliotheek, the Dutch National Library. 82 Biogeographers have of course been working on these questions since the foundations of 83the discipline (Box 2). At the coarsest of scales it is what biogeographical regionalization 84schemes are all about (Wallace 1880, Holt et al. 2013). In practice, the placing of distant 85oceanic islands into regionalization schemes has proven problematic because such islands 86often exhibit multiple source regions (Jønsson and Holt 2015). However, efforts have been 87made to draw lines sub-dividing basins. Examples include the subdivision of the South 88East Asian / island region by Wallace’s Line, Weber’s Line and etc., based on 89zoogeographical data (Whittaker and Fernández-Palacios 2007) or of the region 90based on phytogeographical data (e.g., Renvoize 1979). 91 92Box 2. Extract from the preface of The Archipelago, by 93(1869, Vol. I). 94The question of how to treat complex archipelagic regions has been of interest since the 95foundations of the discipline of biogeography. In an attempt to identify distinct units within 96the broad Malay archipelago, Wallace stated: 97 “…I divide the Archipelago into five groups of islands, as follows: I. The Indo-Malay Islands: comprising the 98Malay Peninsula and , , , and , II. The Group: comprising the islands of 99Timor, , , and Lombock, with several smaller ones, III. Celebes: comprising also the Sula 100Islands and Bouton, IV. The Moluccan Group: comprising Bouru, Ceram, Batchian, Gilolo, and Morty; with the 101smaller islands of Ternate, Tidore, Makian, Kaióa, Amboyna, Banda, Goram, and Matabello, V. The Papual 102Group: comprising the great island of , with the Aru Islands, Mysol, Salwatty, Waigiou, and several

3 3 103others. The Ke Islands are described with this group on account of their ethnology, though zoologically and 104geographically they belong to the Moluccas…” 105Later, he referred to the “Philippine Archipelago” as part of : an 106archipelago within an archipelago. 107 108 More recently, efforts have been made to apply sophisticated modern methods of network 109analysis to (the island region between South-East Asian and Australasian 110continental shelfs) and to the Caribbean that do more than simply identify the boundaries 111between different sub-regions (i.e., groups of islands), also called modules in network theory. 112These analyses determine the degree of compositional connectedness based on species 113distributions and identify the degree of local vs regional topological linkage using null 114models to assess the significance of the linkages. In two papers, Carstensen et al. (2012) and 115Dalsgaard et al. (2014) develop this approach to identify four biogeographical roles for 116islands in the network: (i) network hubs are islands possessing both many local species and 117many shared across the region; (ii) module hubs have many local species but few of regional 118distribution; (iii) connectors possess a few local species but many shared across the region; 119and finally (iv) peripheral islands have few local species and few shared regionally. Their 120analyses identified four modules within Wallacea and six within the , assigning 121islands within these modules to the four categories just listed. In general, remote large islands 122tend to possess high richness of endemics and therefore feature local linkage, whereas 123stronger regional topological linkages, reflecting richness of non-endemics, is characteristic 124of typically smaller islands distant from mainland sources but situated near the boundaries 125between modules. These analyses thus help determine, within complex island regions, how 126best to delimit archipelago membership and inter-connectedness from a biogeographical 127perspective. 128 In slightly simpler circumstances than Wallacea, the Caribbean, or the Indian Ocean, the 129North Atlantic archipelagos west of Northern and Iberia have been grouped 130phytogeographically into the Macaronesian region (Fig. 2; Vanderpoorten et al. 2007, 131Whittaker and Fernández-Palacios 2007, Torre et al. 2018), a label that has recently been used 132for a new line of gin distilled in the Canaries. Rather fine it is too: evidently benefitting from 133the indigenous botanical ingredients. But, for analytical purposes, should we lump the islands 134at the level of the Macaronesian region, or by archipelago (Canaries, Azores, etc), or even, for 135the Azores, for example, sub-divide the archipelago into three sub-groups? Perhaps the 136answer depends on the question being asked? We should also note that the boundaries and 137even the validity of Macaronesia itself has also been the subject of controversy, with evidence 138to suggest that the Cape Verde islands, far to the south (Fig. 2), may not properly belong in a 139grouping with the other archipelagos (Azores, Madeira, Savage [Salvage] Islands, Canaries) 140and that the degree of Macaronesian distinctiveness depends on the choice of taxa (cf. 141García-Talavera 1999, Fernández-Palacios and Dias 2001, Vanderpoorten et al. 2007). 142

4 4 143 144 145Figure 2. Macaronesia is a recognized but controversial biogeographical (in origin 146phytogeographical) region consisting of the archipelagos shown, together with a narrow 147coastal strip of north- and with affinities to the tip of the (after 148García-Talavera 1999). 149 150 To accommodate the notion of different degrees of connectivity between and among 151groups of islands, we suggest adoption of the term meta-archipelago (Box 1). This term does 152not yet appear to be in common usage in island biogeography, although some very brief 153mentions have occurred (e.g., see Kueffer et al. 2016, Triantis et al. 2016; the latter defining 154meta-archipelagos as the archipelagos of archipelagos). However, the term has appeared in 155cultural studies, in which it appears to trace back to the work of the Cuban writer Antonio 156Benítez-Rojo (see Benítez-Rojo and Maraniss (1985) or Benítez-Rojo’s (1992) The repeating 157island: the Caribbean and post-modern perspective). In his work the meta-archipelago is 158described as a chaos “having neither a boundary nor a center” and within which culturally, 159each island is a “copy of a different one, founding and refounding ethnological materials like 160a cloud will do with its vapor.” Well, we get the drift and the concept proposed captures much 161of what we are after, particularly in the idea of replication of units which actually embrace

5 5 162difference as well as degrees of connectivity, and the notion of the importance of connection 163not only amongst islands but between the meta-archipelago and other areas. Yet, for 164biogeographical purposes we need a rather different formulation. 165 We propose that the term archipelago be used for a group of islands, typically closely 166spaced, which have historically exchanged biological information and which have continued 167to do so, with significantly higher frequency than they do with any other land masses. Mostly, 168islands within archipelagos have similar origins and geo-environmental dynamics, and share 169a common source pool(s). By contrast, the meta-archipelago should be used for a collection 170of nearby archipelagos whereby the information exchange is at a lower level (see Fig. 3), yet 171has been and remains sufficient to denote the membership of the islands as having shared 172legacies distinguishing them from other collections of islands and/or mainlands. The 173archipelagos within a meta-archipelago are likely to embrace more varied origins, geo- 174dynamics and source pool biases. These usages are thus akin, in terms of compositional 175pattern, to the notions of modules and networks (sensu Newman 2006, Carstensen et al. 2012, 176Poisot 2013, Thébault 2013, Dalsgaard et al. 2014) but have perhaps broader intuitive appeal, 177extendable across historical and contemporary pattern and process, from oceanic island 178systems (in which many species are generated through in situ diversification) to networks of 179habitat islands (Box 1). 180 181

182 183Figure 3. The Meta-archipelago: in this hypothetical case, a constellation of three 184archipelagos that have and continue to exhibit a meaningful level of information exchange 185(e.g., propagules) signified by the lines of varying thickness joining the islands; exchanges 186among constituent archipelagos should occur significantly less often than the level of 187exchange typical within an archipelago, but significantly more than with other areas. Inspired 188by a sketch in Poisot (2013). 189 190 In a previous essay, Triantis et al. (2016) introduced the idea of the meta-archipelago level 191without, however, elaborating on it. In illustration, they mentioned Tarphius beetles, a genus 192with representatives in both Old and New and which are monophyletic at the level of 193Macaronesia (Amorim et al. 2012). They occur on three of the Macaronesian archipelagos 194(Madeira, Azores and Canaries), and are also monophyletic for the Azores as a whole,

6 6 195although on an island level within the Azores this is not the case and instead multiple 196colonization events have been invoked. Such data provide clear indication of more frequent 197exchange amongst islands within an archipelago than between archipelagos, while justifying 198the treatment of the meta-archipelago as being distinct from the mainland source regions. 199Other plant and animal lineages have also radiated repeatedly in multiple archipelagos within 200Macaronesia, while typically generating single island endemic species on particular islands 201(Price et al. 2018). Similar patterns are found in other complex island regions (e.g., French 202Polynesia: Gillespie et al. 2008). 203 It seems likely that the appropriate scales of separation of membership and the particular 204data and metrics best suited to identify groups of islands as belonging to either archipelagos 205or meta-archipelagoes will depend on the particular biogeographical purposes. Depending on 206the data that are fed into such analyses and the methods of analysis selected, it may be 207possible to develop these approaches to emphasize either contemporary patterns of movement 208and exchange, or to emphasize past process regimes, reflecting deeper time evolutionary 209linkages or, for example, the regimes of currents, climate and sea-level conditions of the 210Pleistocene glacial episodes (Norder et al. 2018). For at least some of these purposes, the 211analytical tools are already well developed (e.g., Dalsgaard et al. 2014, Torre et al. 2018, 212Triantis et al. 2018). 213 In some geographical circumstances (e.g., islands within long, thin lakes), islands may 214actually be exchanging biological information with the mainland more frequently than with 215each other. In such circumstances, the groups of islands concerned may be useful for many 216island biogeographical purposes, but arguably do not warrant the label of archipelago by the 217above definition. Paradoxically, by this approach might the islands of the Aegean Sea be 218deemed merely an island group or region, rather than one archipelago, or meta-archipelago? 219Here the evidence for five out of the nine taxa considered in Triantis et al. (2018, p. 287), is 220that, for example, Crete and the surrounding islets are quite distinct from the rest of the 221Aegean islands, and thus Crete and its islets can be seen as an archipelago within an Aegean 222meta-archipelago. 223 Were it the case that all taxa have similar scales of interaction with fragmented land- and 224seascapes, then it might be realistic to think of prescribing a single framework of islands, 225archipelagos and meta-archipelagos. But as previous regionalization and filter effect analyses 226have shown, and as recent modularity analyses of Macaronesia also show (Torre et al. 2018), 227this is not the case. Hence, the distributions and exchanges of more vagile taxa frequently 228span multiple archipelagos, whilst the least vagile taxa exhibit largely within-archipelago or 229within-island scales of distribution and exchange. Even within a single taxon (e.g., 230bryophytes, seed plants, beetles, birds, land mammals, etc), there is always a significant span 231of movement or dispersal attainment. In using the terms island, archipelago and meta- 232archipelago, therefore, there is a further question of how broadly applicable across different 233taxa the labelling needs to be for it to be useful? Methods and approaches applied for 234biogeographical regionalization and/or network analyses that use distributional (sometimes 235with phylogenetic) data from multiple taxa might be of use in resolving this question (e.g., 236Holt et al. 2013, Ficetola et al. 2017). 237 238 Do we need a new term? The proliferation of terms is not always helpful to a discipline, 239although arguably the greater problem is the inconsistent use of the terms we have. In this 240instance, we suggest that the term meta-archipelago has appeal and may prove useful. In 241particular, it may encourage ecological island biogeographers to conceptualize the islands 242they study as belonging not simply to one group, framed in relation to a particular, distant 243mainland, but to consider and to analyse how the biota of each island may be part of a loose,

7 7 244but structured network of concentric layers of relatedness and exchange. The layers of 245archipelago and meta-archipelago provide the two closest layers of the networks that exist 246around many islands, while the failure to identify such patterns of linkage would identify an 247island as either a truly isolated island, an island that sits in the pocket of a dominant 248mainland, or one that belongs to a continuum or patchwork of more or less connected habitat 249patches. Ecologists and biogeographers are familiar with the terms meta-population and 250meta-community, which denote the subdivision of populations and communities into areas 251that are insufficiently connected to form a single entity but yet are not entirely independent of 252one another (e.g., Leibold and Chase 2017). The term meta-archipelago in its essence, simply 253extends this concept into island biogeographical pattern and process, embracing both 254ecological-island-biogeography and ‘evolutionary/historical-island biogeography’, hopefully 255encouraging a free flow of discussion bridging these traditions. 256 257------258References

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337Submitted: 29 October 2018 338First decision: 20 November 2018 339Accepted: 30 November 2018 340Edited by Michael N Dawson

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